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Asteraceae) in Southern Africa S.Afr.1.Bot., 1991, 57(6): 325 - 330 325 Pollen morphology and its taxonomic value in Brachylaena (Asteraceae) in southern Africa S.S. Cilliers Department of Plant Sciences, Potchefstroom University for CHE, Potchefstroom, 2520 Republic of South Africa Accepted 21 August 1991 The pollen morphology of all the southern African representatives of Brachylaena R. Sr. was investigated using light and scanning electron microscopy. On the basis of pollen morphology it is possible to recognize two groups in the genus. B. discolor (including B. unifloral. B. elliptica, B. glabra, B. neriifolia and B. rotundata are included in Group I, while B. huillensis and B. ilicifolia are the only representatives of Group II. The differ­ ences between the groups primarily concern length and width of colpi, shape of colpus ends, length and width of spinulae, frequency of spinulae, finer tectum sculpture and exine stratification. The possible taxo­ nomic significance of pollen morphology is briefly discussed. Die stuifmeelmorfologie van al die suider-Afrikaanse verteenwoordigers van Brachylaena R. Br. is met behulp van lig- en aftas-elektronmikroskopie ondersoek. Op grond van stuifmeelmorfologie kan twee groepe binne die genus onderskei word. B. discolor (insluitend B. unifloral. B. elliptica, B. glabra, B. neriifolia en B. rotun­ data ressorteer in Groep I, terwyl B. huillensis en B. ilicifolia die enigste verteenwoordigers van Groep II is . Die belangrikste verskille tussen die twee groepe is ten opsigte van lengte en breedte van kolpusse, vorm van die kolpuspunte, lengte en breedte van spinulas, frekwensie van spinulas, fyner skulptuur van die tektum en eksienstratifikasie. Die moontlike taksonomiese waarde van stuifmeelmorfologie word kortliks bespreek. Keywords: Asteraceae, Brachylaena, pollen morphology. Introduction Tarchonanthus species. The sexine of the pollen of this type Thorough palynological studies on certain representatives of is 1.6 J1.m thick and consists of two layers of baculae (Leins, the Asteraceae have been carried out by various authors 1971). The nexine is 0.8 - 1 J1.m thick and consists of two [Wodehouse (1935) on external morphology and Stix (1960) parts (nexine 1 and 2). He also mentioned that these pollen on internal morphology] . These studies usually concentrated grains are tricolporate and possess narrow colpi with pointed on taxonomically useful characters. Since the usefulness of ends, while the tectum is covered with spinulae. The B. huil­ pollen as a taxonomic tool is restricted by the number of lensis pollen type was found in B. huillensis and B. ilicifolia characters that can be observed by means of light micros­ only and the sexine of their grains is 0.7 J1.m thick and copy (LM) (Wodehouse, 1928), it became apparent to tax­ consists of a single bacula layer. The nexine is undifferen­ onomists that additional pollen characters would have to be tiated and is 0.5 J1.m thick. These pollen grains are tri- or obtained by using transmission (rEM) and scanning (SEM) tetracolporate and possess much wider colpi with rounded electron microscopes (Skvarla et al., 1977). ends and the tectum is also covered by spinulae. The Bra­ As early as 1966 Skvarla and Turner reported on TEM chylaena ramif10ra pollen type is unique to this Madagascar studies of pollen of representatives of most of the tribes of species. the Asteraceae, while Clark et al. (1980), Tomb et al. Leins (1971) also questioned the validity of the inclusion (1974) and others did SEM studies on a selection of repre­ of the subtribe Tarchonantheae in the tribe Inuleae, on the sentatives of the family. In spite of the palynological basis of their pollen morphology. He recommended the clas­ research in the family, relatively few studies on comparative sification of this subtribe in the tribe Mutisieae, while pollen morphology of taxa were carried out at infrageneric Skvarla et al. (1977) suggested its inel usion within the tribe level (Keeley & Jones, 1977). Anthemidae, also based on pollen morphology. Brachylaena To date, pollen of Brachylaena R. Br. has been studied by was also placed in three other tribes over the years. Keeley means of LM only (Erdtman 1966; Leins 1971). The pollen and Jansen (1991), however, recognized a new tribe, the of Brachylaena is tricolporate and subprolate with an exine Tarchonantheae, with Brachylaena and Tarchonanthus the stratification similar to that of Tarchonanthus (Erdtman, only members, on the basis of chloroplast DNA variation. 1966). Material from one locality only of each of Brachy­ Unfortunately, Leins (1971) did not study pollen of all the laena discolor, B. rotundata and B. neriifolia were studied southern African species of the genus. He also ignored some by Erdtman (1966). representatives of the taxonomically problematic B. dis­ Based on LO analyses of the pollen of most of the mem­ color-unif1ora species complex and in five cases he based bers of the tribe Inuleae, Leins (1971) distinguished three his results on pollen from only one specimen. pollen types in the subtribe Tarchonantheae (to which Bra­ To determine whether pollen morphological characters chylaena belongs), mainly on the basis of exine stratifica­ have taxonomic value amongst the southern African repre­ tion. The Tarchonanthus camphoratus pollen type was sentatives of Brachylaena, it is essential to gain information found by Leins (1971) in four Brachylaena species (B. dis­ from specimens of all the taxa collected from various color, B. rotundata, B. neriifolia and B. elliptica) and four localities. The aim of this study was to search for additional 326 S.-Afr.Tydskr.Plantk., 1991, 57(6) taxonomically useful characters by using the SEM. To check tectum sculpture. Initially the pollen grains were cleaned by whether the results of Leins (1971) on exine stratification the acetolysis method of Erdtman (1960). Because of colpar hold true for all the Brachylaena taxa in southern Africa, the damage and the fact that this method is time-consuming, SEM studies were complemented with LM studies. the filter technique of Bredenkamp and Hamilton-Attwell (1988) was subsequently used. Materials and Methods The polar axis (P), equatorial width (E) and the thickness Pollen of all the southern African taxa of Brachylaena of the exine and its parts were measured by means of an LM [B. uniflora is included in B. discolor subsp. transvaalensis, provided with an eye piece micrometer. All measurements (Ci II iers , 1990)] were studied by means of LM and SEM. were based on a minimum of ten pollen grains per speci­ Pollen from herbarium specimens from a minimum of four men. The frequency of spinulae was determined from an localities each were studied (Table 1). average of three counts in an area of 2 f-Lm x 2 f-Lm (4 Data on the following characters were recorded: f-Lm 2) in the mesocolpium, by using SEM prints of similar (a) Polarity, size and symmetry of pollen grains (SEM and magnification. LM). The descriptive terminology used mainly follows Erdtman (b) Shape, as well as the amb: equatorial diameter (SEM (1969) and the attempts at standardization offered by and LM); polar diameter (SEM and LM). Reitsma (1970) and Nilson and Muller (1978). The shape of pollen grains in equatorial view was determined from the (c) Aperture type (SEM and LM). shape classes of Erdtman (1966), while terms proposed by (d) Colpus structure (SEM): length of colpus, in equatorial Faegri and Iverson (1975) were used to describe the shape view; width of colpus, in equatorial view; shape of of pollen grains in polar view. colpus ends, in polar view; surface area of the apo­ colpium. Results (e) Tectum sculpture (SEM): length of spinulae; frequency of spinulae in the mesocolpium; finer tectum sculpture Polarity, size and symmetry of the pollen grains (presence of other structures, except spinulae). The pollen grains of all the taxa are released as monads and (f) Exine stratification (LM). are radially symmetrical and isopolar. The polar axis (P) and Some preparation techniques may influence the size and equatorial width (E) of the pollen grains of B. huillensis shape of the pollen grains of representatives of the Aster­ (P = 30 - 37 f-Lm and E = 26 - 35 f-Lm) are larger than those aceae (Cuzma Velari, 1984) and therefore untreated grains of all the other species (P = 26 - 34 f-Lm and E = 20 - 30 were used. Pollen was obtained from unopened florets, f-Lm) . However, as a result of the overlap that exists in the respective dimensions, it is not possible to distinguish be­ mounted directly onto stubs using a carbon glue (,Leit-C tween them on the basis of these characters only. The length nach Gocke', Neubauer Chemikalien), spuller-coated with of the polar axis of the pollen grains of all the Brachylaena gold/palladium and viewed using a Cambridge Stereoscan taxa studied, is 25 - 50 f-Lm, the grains therefore being 250S SEM. Pollen from the same specimens was also pre­ medium in size (ME). pared for LM studies following the method of Coetzee and Van der Schijff (1979). Although only the shape and size of Shape of pollen grains the pollen grains could be studied from these preparations, the pollen was also cleaned to study col pus structure and The shape of the pollen grains of most of the taxa is subprolate (Figure 1), and occasionally prolate spheroidal. Most of the pollen grains of B. huillensis are prolate Table 1 Specimens used in the pollen morphological spheroidal (Figure 2), but sometimes oblate spheroidal or study subprolate. Since the variation in pollen grain shape is continuous, it is not possible to distinguish B. huillensis Taxon Collector, number and herbarium from other species on the basis of their pollen grain shapes. B. discolor subsp. Cilliers 123, 135 (pUC), Hilliard 1673 (UN), Law The pollen grains of most of the taxa studied appeared discolor 5 (NBG), Taylor 206 (UN), Thode Al532 (NH), Venter 1927 (ZULU).
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